Journal of Global Biosciences: Research Paper

Journal of Global Biosciences
ISSN 2320-1355
Volume 6, Number 4, 2017, pp. 4876-4895
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Research Paper
EFFECTS OF ORGANIC MANURE ON SOIL PROPERTIES AND HEALTH
AND GROWTH PERFORMANCE OF CACAO (Theobroma cacao L) IN
SOUTHWESTERN NIGERIA
Kayode B. Adejobi1, Samuel O. Agele2, and Peter O. Aiyelari2
1Agronomy Section,
Cocoa Research Institute of Nigeria,
PMB 5244, Ibadan, Nigeria.
2Department of Crop,
Soil and Pest Management,

Federal University of Technology Akure,
Nigeria.
Abstract
Soil fertility status and activities of soil borne microbes are major factors
influencing cacao growth and establishment on the field; in this study, the
manurial potentials of some organic wastes and their effects on soil
health and growth of cacao in Ibadan and Owena soils of Southwestern
Nigeria were investigated in 2012. The experiments were Randomized
Complete Block Design (RCBD) with 3 replications. The fertilizer types
were: Goat Dung (GD), Sunshine Organic and Organo-Mineral Fertilizers
(OF and OMF) and NPK (15:15:15) which were applied at 0, 200, 400 and
600 kg/ha. Plantain suckers were established at 3 x 3 m in June, 2011 as
shade crop and cacao seedlings were transplanted into the plots one year
later (June, 2012). The fertilizers were applied to treatment plots one
month after cacao seedlings were transplanted using ring method. Data
were collected on cacao growth parameters such as plant height, stem
diameter, number of leaves and number of branches, monthly
commencing at 3 months after transplanting (MAT) and for 96 weeks
after transplanting (WAT). Biological properties (arbuscular myccorhizal
spores, nematodes, moulds, yeast and bacteria) were also determined on
the soil samples using standard procedures. Goat dung (GD), organo-
mineral fertilizer (OMF) and organic fertilizers (OF) had pH ranged
between 7.00 and 8.17 while they contained in similar order 2.86, 0.63
and 3.64 g/kg organic carbon (OC), 4.80, 1.09 and 6.27 % organic matter
(OM), 1.26, 0.06 and 2.16% Nitrogen (N), 113.24, 138.06 and 7.08
cmol/kg Phosphorus (P), 2.60, 2.00 and 13.10 cmol/kg Calcium (Ca)
respectively. All fertilizer types and rates at Ibadan and Owena
significantly increased cacao seedlings’ growth relative to the control.
Application of GD, OMF and OF increased albuscular mycorizal spores
Journal of Global Biosciences Vol. 6(4), 2017 pp. 4876-4895
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relative to NPK and the control. The population of other soil microbes
(bacteria yeast and moulds) was also influenced by fertilizer types and
rates. Application of 600 kg/ha NPK significantly reduced soil nematode
population when compared with other fertilizer types, rates and the
control plots in Ibadan, while 400 kg/ha OF and 200 kg/ha OMF
significantly reduced nematode population relative to the control in
Owena.
Key words: cacao, fertilizers, biological properties, growth.
INTRODUCTION
Cacao was classified as a member of the family Sterculiaceae until recently when
Alverson et al. (1999) through the application of molecular marking technology,
reclassified the crop as a member of the family Malvaceae. Its natural habitat is the
lower middle storey of the evergreen rain forest. The importance of cocoa in the Nigeria
economy remains high. Cocoa exports have been and will continue to be a significant
factor in the economic growth of Nigeria. For instance, in 1969, cocoa alone earned
N106 million which accounted for 40% of all agricultural exports for the year (Federal
Office of Statistics 1972).
Despite negative effect of government policy, cocoa remains the highest foreign
exchange earner of all agricultural export crops which the Structural Adjustment
Programme (SAP) sought to promote as a development policy objective in Nigeria
(Tijani et al. 2001). However, the discovery of crude oil in large quantities has brought a
downward trend in Nigeria cocoa production and position in the world market (Ayoola
et al. 2000). Côte d’lvoire, which was third in Africa with 143,000 tonnes behind
Nigeria’s l96, 000 tonnes in 1970, is now the largest producer in the world with 1.2
million tonnes annually accounting for about 40 % of total world production. Nigeria is
now the fourth largest producer after Côte d’lvoire, Ghana and Indonesia (ICCO 2005).
The practice of planting cacao seedlings under banana or plantain widely preferred with
increase risk of Meloidogyne incognita infection on cacao seedlings, as the nematode is
frequently associated with the shade crop and parasitic nematodes have become
common pests of cacao in West Africa (Fademi et al. 2006). Plant-parasitic nematodes
as pests, pose threat to young cacao seedlings at both establishment and fruiting stages
of cocoa. Meloidogyne incognita appears to be the most common nematode of cacao in
Nigeria. The symptoms observed were, die back, stunting, wilting, chlorosis, reduction
in the size of the leaves and galling of the root or complete death of the seedlings
(Orisajo and Dongo 2005). Hence there is an urgent need to develop new management
tools that are affordable, accessible, environmentally and toxicologically safe in
controlling the pests. Conveniently, the use of resistant planting materials is the most
economic approach to nematode control in plants. However, cacao was not listed among
the eight nut/fruit trees which have locally available nematode – resistant or nematode-
tolerant seeds or rootstocks (Sasser and Frekman 1987) Nematodes, hence, pose a
threat to cacao that can not be addressed through resistance or tolerance of planting
materials (Sasser and Frekman 1987). Therefore other methods of control have to be
investigated. Furthermore, it is well documented that the increase in plant growth
resulting from Arbuscular Mycorrhizal Fungus (AMF) association is usually due to
increased mineral elements uptake by the hyphae from the soil and increased uptake of
macro-elements other than P has also been reported (Hodge et al., 2001) as well as
increased uptake of some micro-elements (Kothari et al. 1991a). In addition
Mycorrhizae have been shown to play an important role in maintaining soil physical
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properties (Degens et al. 1996). It has also been observed that soil organic amendments
foster beneficial soil micro organisms (Doran 1995). Limited field studies have been
conducted to determine the impact of soil amendments on microbial communities in
organic and conventional production systems (Gunapala and Scow 1998). However, it
has been shown that microbial activity and biomass is higher in fields when organic
manures are applied compared with application of synthetic fertilizers (Doran 1995).
The aim of this study therefore is to examine the influence of fertilizers (organic and
inorganic) on the diversity of soil microbes, soil chemical properties and growth of
cacao seedlings on the field.
MATERIALS AND METHODS

Field experiments were conducted at Ibadan, headquarters of the Cocoa Research
Institute of Nigeria (CRIN), and in Owena, a CRIN Sub-station in Ondo State.
Ibadan experimental location
Ibadan, Oyo State, is located on latitude 070 10' N and longitude 030 52’E, and an
altitude of about 122 meters above sea level in the humid tropical rain forest zone of
Nigeria. Ibadan is characteristically hot and humid, reputed for seasonal rainfall, high
temperatures and high humidity and distinct dry and rainy seasons. The dry season
extends from early November to March and is characterized by dry atmosphere and
intense scorching sun. The rainy season which is characterized by high humidity and
cloudy sky, runs from end of March to early November. There is usually a two-week dry
spell in August. The rainy season has an average of 8 rainy days in a month at the
commencement of the rains, this increase from April to about 15 rainy days in
September. The annual rainfall is between 1200-1500 mm. The maximum temperature
ranges between 26 to 35 o C with an average of about 30.1oC while the minimum
temperature ranges from 15 to 24o C with an average of 19.5oC. Relative humidity is
high throughout the year and ranges from of 50 to 89 % with an average of 79 %. There
are seasonal variations in the values of relative humidity, which varies from 65 to 89 %
during the rainy season and 46 – 70 % during the dry season.
Owena experimental location
The Owena Substation of the cocoa Research Institute of Nigeria, located in Ondo State,
lies on latitude 07o N and longitude 05o, 71E and at an altitude of about 22.5 meters
above sea level. Owena is situated at about 21 kilometres south–east of Akure between
Akure and Ondo. There are two seasons: rainy (wet) season which spans late March to
November of the same year and a dry season that runs from December of one year to
late March of the following year. There are 15 to 17 rainy days per month during the
rainy season. The dry season on the contrary to the situation in Ibadan` has
characteristic of scanty rainfall, the average ranges between 1500 – 1600 mm per
annum. The maximum temperature in Owena is usually between 28 and 34o C while the
daily minimum temperature ranges between 18 and 23o C. The relative humidity during
the rainy season ranges from 69 to 80 %, and between 56 and 64 % during the dry
season.
Acquisition and preparation of experimental materials
Seedlings cacao F3 Amazon genotype was collected from CRIN Seed Garden, while
plantain suckers were collected from experimental plots in both Ibadan and Owena.
Experimental plots of about 30 by 120 m were mapped out and the experiment was laid
out in rows of 3 x 3 m. Sunshine organic and organo-minerals fertilizers used for the
experiments were obtained from the Ministry of Agriculture, Ondo State, while the
N.P.K. 15:15:15 was obtained at Ayedaade Local Government Gbongan, Osun State. Goat
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dung manure was obtained from Ilesha Garage Akure, Ondo State. The goat dung was
collected dried and carefully sorted to remove foreign materials and packed in 50 kg
bags for application on the field.
Treatments and experimental design
Field experiments were conducted in two stations of the Cocoa Research Institute of
Nigeria (CRIN), Ibadan Headquarters and Owena Sub-Station, Ondo State. The
experiments were conducted between June, 2012 and June, 2014. The experiments
were Randomized Complete Block Design (RCBD) with 3 replicates. The 4 fertilizers
were: Goat Dung (GD), Sunshine Organic and Organo-Mineral Fertilizers (OF and OMF)
and NPK 15:15:15 and the rates of application were: 0, 200, 400 and 600 kg/ha. Lay-out
of each experimental site (Measurement, pegging and holing) was carried out before
planting. Four hundred and thirty two (432) plantain suckers were planted at the
spacing of 3 x 3 m in each of the experimental sites in Ibadan and Owena between
second and third week of June, 2011, while Four hundred and thirty two (432) five
months old cacao seedlings (F3 Amazon) of average height of 46 cm (already raised in
the nursery) were transplanted in to one year old plantains in 2012 at the spacing of 3 x
3 m on each of the sites (Ibadan and Owena). Four plants were randomly tagged for data
collection. Top soil samples were collected randomly from each experimental sites
(Ibadan and Owena) using soil auger. The samples were bulked and air dried before
being subjected to routine laboratory analysis of particle size analysis was determined
by the hydrometer method (Kettler et al. 2001) and organic cabon content (OC) by the
potassium dichromate oxidation method (Zhang et al. 2001). Soil pH was read on pH
meter (1:1 water). Organic matter was determined by the Murphy blue coloration and
determined on a spectronic 20 at 882um (Murphy and Riley, 1962). Soil potassium (K),
calcium (Ca) and Magnesium (Mg) were extracted with lMNH4 OAC, PH7 and were
determined with flame photometer; Mg was determined with an atomic absorption
spectrophotometer. The total nitrogen (N) was determined by the Microkjedahl method
(AOAC, 1990). Two grammes (2 g) each of the organic fertilizers used were also
analyzed for nutrient composition. The fertilizers were applied to treatment plots one
month after transplanting using ring method of fertilizer application at 5cm away from
the base of cacao (July, 2012). Monthly Data collection on growth parameters (plant
height, stem diameter, number of leaves and number of branches) commenced 3
months after transplanting. The experiments were monitored for 24 months (96 weeks
after planting). At 15 months after transplanting, soil samples were collected from
treatment plots and were processed and analyzed for and soil biological properties
(Arbuscular myccorhizal colonization, soil nematodes, moulds, yeasts and bacteria)
were determined using standard procedures.
Data were collected on the growth parameters of cacao seedlings such as: Plant height
measured in centimetre using a meter rule on the surface and the tip of the main stem;
Number of leaves was counted. Stem diameter was measured in centimetre with the use
of Vernier’Caliper 30 cm above the ground level. Number of branches was also counted.
These growth parameters were taken monthly for 24 months commencing from 3
months after transplanting (3 MAT).
Soil samples collection and analysis

Top soil samples were collected randomly at the depth of 15 cm from each of the
experimental sites at both locations (Ibadan and Owena) with the aid of soil auger at
30cm depth. For the pre-cropping analysis, the samples were bulked together and
mixed thoroughly, air dried at room temperature and analysed for various elements.
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Particle analysis was carried out using the hydrometer method as described by
Bouyoucos (1951). The pH was determined in water (1:1 soil: water ratio) using a pH
meter with glass electrode as described by Jackson (1965). Total Nitrogen was
determined using kjeldahi procedure as described by Jackson (1965). Organic carbon
content was determined using the Walkey-Black method (Nelson and Sommers 1982).
Phosphorus determination was done by the Bray method as described by Bray and Kurt
(1945).
Exchangeable K, Ca, Mg and Na were determined by extraction with ammonium acetate

and the amounts of K, Ca and Na in the filtrate were determined using flame photometer
with appropriate filter while Mg was determined using a perking Elmer Atomic
Absorption Spectro photometer (AAS).
Determination of diversity of soil microbes
Soil samples were collected from 15 cm depth in each experimental plot around cacao
seedlings, placed in nylon bags, thoroughly mixed, and air-dried for 48 hours. The
samples were sieved through 2 mm sieve and stored in the shade at room temperature
until they were analyzed.
Soil assay for nematodes
The soil was assayed to confirm the presence and the initial population density of the
nematodes. A 100g sub-sample soil was assayed for nematodes using the White-head
and Hemming (1965) tray modification of Baermann (1917) technique as described
below:
One hundred grams soil was put into a set up that has two plastic sieves with extractor
tissue sandwiched in between. The plastic sieves with the soil were thereafter placed in
a plastic bowl, and water was added to the extraction bowl just enough to wet the soil.
The set-up was left undisturbed for 48 hours. Thereafter, the plastic sieve containing
the soil was removed briskly, and the nematode suspension in the bowl was poured into
a nalgene wash bottle and allowed to settle (Caveness 1975). The supernatant was
siphoned out, and the suspension containing nematodes was then poured into a labelled
beaker, and adjusted to 10 ml by adding water. This was homogenized and 1ml of the
suspension was taken with the use of pipette, dispensed into the Doncaster (1962)
nematode counting dish and examined under a stereomicroscope. Nematodes was
transferred with a sharpened broom stick to a slide with a drop of water, covered (with
a cover slip) and examined under a compound microscope with a 40, 60 and 100 X
objective for identification using taxonomic keys (Luc and de Guiran 1990) and counted.
The identification and counting were repeated three times and mean population of
nematodes per sample were calculated. The various procedures followed for soil
assayed were outlined by Coyne et al. (2007).
Extraction of vesicular arbuscular mycorrhiza (VAM) fungal spores from soil
The wet-sieving method (Gerdamann and Nicholson 1963) was employed. A 100 g sub-
sample soil was suspended in 750 ml tap water in 1000 ml capacity plastic container for
sedimentation after which the suspension was mixed vigorously. The soil suspension
was strained through four sieves of 710, 500, 250 and 53µm mesh sizes (Model
Endecotts UK) arranged in the order, 710, 500, 250 µm, to remove rock fragments and
coarse woody and root debris. The soil suspension was finally strained through sieve 53
µm to collect the spores mixed with the solid matter; thereafter the solid matter
collected was transferred to 50 ml centrifuge tube. Water was added to the tube and the
soil sample was re-suspended. The tubes were centrifuged at 1800 rpm for 5 minutes.
The supernatant which contained floating organic materials including dead spores was
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discarded. The pellet was re-suspended in sucrose solution (440 g/l). Carefully, the
tubes were balanced and centrifuged up to 1800 rpm for 2 minutes and stopped
immediately. Rapidly, the supernatant was sieved (32 micron) and washed thoroughly
(at least 1minute) to replace the sucrose and alleviate osmotic stress on spores. The
pellet left in centrifuge tube was discarded. Carefully, all of the solid materials from the
sieve were washed into the petri dish marked with a grid to facilitate spore counting
and collection under a stereo- zoom microscope. The spores in suspension were filtered,
counted and identified using Stereo-zoom microscope (Model Leica MZ125) at
magnification 40 X and Compound microscope
Enumeration of Bacteria, Yeast and Moulds in the soil in soil sample
Nutrient agar (NA) and Potato Dextrose Agar (PDA) were prepared according to
manufacturers specification (BIOMARK) by dissolving 28 g of NA in a litre of distilled
water in a conical flask while 39 g of PDA was dispersed in a litre of distilled water
separately, allowed to soak for few minutes, thoroughly mixed, cotton plugged, covered
with aluminium foil and then sterilised by autoclaving at 15 lbs pressure (121o C) for 15
minutes. The media were allowed to cool and mixed well before dispensing. The
standard procedures for determining the total number of soil microbes were adopted
for bacteria, yeast and moulds culturing. Suspension of the soil samples was prepared
with sterile distils water and serial dilution of 5-7 factors was made for accurate
counting. Then 1ml portion of the sample was aseptically pipetted into different sterile
Petri dishes with the aid of sterile needle and syringe. About 20ml of the cool (40o C)
sterile molten agar media was added, swirled gently for even distribution of inoculums
within the plate, then allowed to set and incubated. Nutrient agar plates for bacteria
were incubated at 37oC for 24hrs while PDA plates for fungi and yeast were incubated at
27o C for 48 – 72 hrs. At the end of incubation period, the bacteria that grew into
colonies were counted using colouring counters while yeast and fungi colonies were
squarely counted and recorded appropriately per gram of the soil samples investigated
respectively.
STATISTICAL ANALYSES
Analysis of variance was performed on all data to test the treatment effect on different
parameters and significant means were separated using Tukey’s Honest Significant
Difference (HSD) (P < 0.05).
RESULTS AND DISCUSSION

The pre- planting soil physical and chemical properties of the organic wastes used for
the experiments are presented in Tables 1 and 2.The results of the particle size analysis
of Ibadan and Owena soils showed that the soils were sandy loam and Alfisols (Soil
Survey Staff 1999). The silt + clay contents of the soils at Ibadan and Owena (24.98 %
and 25.96 %) respectively were below the 32 % estimated as adequate soils ideal for
tree crops especially cacao (Egbe et al. 1989). Based on the established critical levels
for soils in south western Nigeria, the soils at Ibadan and Owena were acidic with pH
ranging between 4.56 – 5.70 and low in organic matter (1.26 – 1.61 %) compared to the
reported critical levels of 3 % organic matter (Agboola and Corey 1973). The total
nitrogen of Owena soil was less than 0.15 % which is considered optimal for most crops
including cacao and the soil also had low CEC (Ogunwale et al. 2002). This suggests the
need to improve on the soil organic matter and hence the CEC for enhanced nutrient
retention and the release of same to crops upon external fertilizer application (Agboola
and Omueti 1982). The improvement of soil organic matter (SOM) can be achieved by
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organic fertilizer application, either as sole or in combination with inorganic fertilizers

as organo-mineral fertilizer (OMF). The application of inorganic fertilizer to a soil with
low organic matter content is a waste of resources and time (Agboola and Obigbesan
1975). Hence there is need for proper SOM management on Ibadan and Owena soils to
reduce the deleterious effects on soil physical, chemical and biological properties.
Although, available P was also low in Ibadan, this level of available P is considered
inadequate for cacao (Wessel 1971; Egbe et al. 1989). Only Ibadan soil gave
exchangeable potassium above the critical value of 0.3 cmol/kg required for cacao. The
exchangeable Ca+ of Owena soil fell below the critical value of 5 cmol/kg required for
cacao growth. At both locations, the exchangeable Mg+ was adequate for cacao
production. Obatolu (1991) earlier observed the general low Mg+ nutrient contents of
these soils. The low nutrient contents of the soils implied the need for external input of
nutrients in order to meet the requirements for optimal cacao growth. It is obvious that
the soils of both Ibadan and were inherently low in fertility and were therefore
expected to show positive response to soil amendment. The insufficient levels of the
major nutrients in the soils in both locations showed that the soils were depleted in
nutrients and would not be able to meet the nutritional needs of cacao plants unless
external nutrients supply is made to support optimum growth of cacao plants.
Among the organic fertilizers applied, goat dung (GD) produced the highest pH, though
the organic and organo-mineral fertilizers had pH above 5 (acidity levels) which
indicated that they could be effective as liming materials. The organo-mineral fertilizer
(OMF) had the highest available P followed by GD and organic fertilizer (OF) produced
highest percentage N. The results were in agreement with the works of Adejobi et al
(2011a) who reported that GD, OMF and OF were as effective as NPK fertilizer as
sources of plant nutrients. In particular, OF had the highest OM, K, Mg, Ca and Na
concentrations relative to others, this implied fact that OF could be a good source of
these nutrients for plant growth. The organic manure types and rates applied increased
the abundance of Verscular Abuscular Mychorizal Spores (VAMS) in the soil relative to
NPK rates and the control in both Ibadan and Owena (Table 3). The presence or
availability of mychorizal fungi is necessary for proper growth of seedlings in forestry
nurseries and for successful start off on the field after transplanting (Mohammand et al.
2003). This result implied that organic sources of nutrients such as farmyard manure
(FYM), compost, crop residues and slow release mineral fertilizers do not suppress
AMF but stimulate them (Alloush and Clark 2001). However, it has been observed that
over use of organic fertilizers especially those high in P, such as poultry manure, may
impact negatively on AMF. However, the precise effect of organic amendments is
unpredictable on any given soil or with any particular amendment (Alloush and Clack
2001). Jordan, (2000) found that AMF in an organically manured soil were effective at
increasing crop available P comparable to application of super-phosphate. However,
this does not always translate into higher yields even when phosphorous use efficiency
is higher (Galvez et al. 2001). At Ibadan, application of 400 kg/ha OF enhanced
abundance of AMF than other fertilizer types, rates and the control. The lower number
of AMF count recorded under NPK rates was in agreement with the findings of Burrow
and Pfeger (2002) who stated that use of readily soluble fertilizers, particularly N
fertilizer, had negative impact on AMF colonization, population and diversity. Again, the
result of this study showed that plots amended with the fertilizers suppressed the
population of soil nematodes that are associated with cacao in both Ibadan and Owena
experiments (Table 4). Soil treated with NPK fertilizer at 600 kg/ha and organo-mineral
fertilizer (OMF) at 200 kg/ha gave the most effective nematode control at Ibadan and
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Owena experiments. In both Ibadan and Owena soils, highest nematode populations
were recorded on the control plots. Several studies have shown the advantages of using
organic manures in controlling plant parasitic nematodes (Dias et al. 2000; Nagesh 2002
and Salgado et al. 2003). This study has shown that fertilizers like GD, OF, OMF and NPK
have nematicidal effects on soil parasitic nematodes: Meloidogyne incognita, Heterodera
sahatti, Ditylenchus dispsaci, Paralongidorus sali, and Psilenchus sp identified in this
study. This result is consistent with the report of Riegel and Noe (2000) that application
of poultry litter 14 days before planting was optimal for reduction of nematode
population densities. Many factors could affect the response of nematode communities
to nutrient sources. Most importantly, nematode communities were often affected by
the nutrient composition, particularly the C: N ratio of the organic amendments (Ferris
and Matute 2003; Yeates and Boag 2004). In general, amending the soil with organic
materials having low C: N ratio (less than 1:20) resulted in an abundance and
enrichment of opportunist antagonistic microbes (Ferris and Matute 2003; Wang et al.
2004 and 2006) and rapid mineralization of N in the form of NH 4+ or NO3 – for
absorption and uptake by plant roots (Powers and Mcsorley 2000). The fertilizers used
in these experiments have low C: N (1:4, 1:1, 1:6) and this appeared to have resulted in
the suppression of nematode population on cacao seedlings. It has been established that
organic amendments released some chemicals into the soil that are directly responsible
for nematode control. Ricin, a proprein derived from castor bean has nemato-toxic
potential (Rich et al. 1998). The neem tree (Azadirachta indica) contains a group of
chemicals known as limonoids and these compounds have proven highly effective in
nematode control. Phenols and tannins are nematicidal at certain concentrations, and
since some organic amendments added to soil contain high levels of these compounds,
they may have a direct effect on nematode mortality (Badra and Eligindi 1979).
Differences in soil microbial counts (Bacteria, Yeasts and Moulds) as obtained under the
types and rates of fertilizers applied suggest their use for enhancement of fertility and
crop yield (Table 5). The results showed that GD applied at 200 and 600 kg/ha NPK
increased soil bacteria count significantly at both Ibadan and Owena 2012 experiments,
400 kg/ha OMF, 600 kg/ha OF and 200 kg/ha GD significantly enhanced yeast
population at both Ibadan and Owena relative to the control and 200 kg/ha NPK and
400 kg/ha OF increased soil fungi at Ibadan and Owena experiments relative to other
fertilizer treatments and the control. Stimulating effects of fertilizers on soil microbes
was observed in the locations when compared with the control.
The enhancement of soil biological parameters by application of organic fertilizers on
soil at 15 MAT has been reported by other authors (Bian et al. 2008; Emitsev et al.
2010). Additionally, as compared to other fertilizers and the control, the fungal
populations at both Ibadan and Owena were significantly enhanced when 200 kg/ha
NPK and 400 kg/ha OF were applied. The decline in the population of this group of
micro-organisms in the control plots was an expected occurrence (Table 5). The lower
fertilizer rates (200 and 400 kg/ha,) stimulated the development of the bacteria, yeasts
and moulds in the two locations with the exception of bacteria in Owena (Table 5). This
finding was in agreement with the results of Barabas et al. (2002), who reported an
increase in the count and diversity of bacteria, actinomycete and fungal species under
lower mineral nitrogen application. The decreased rates of application of mineral
fertilizer (200 kg/ha) led to increase in the count of the fungal populations at both
Ibadan and Owena. This tendency is frequently associated with changes in the soil
physical and chemical characteristics (Stark et al., 2007) as well as with the alterations
in the structure of soil microbial cenosis expressed through the predominance of
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toxinogenic and phytopathogenic fungi. Also, Barabasz et al. (2002) has cautioned that
inadequate application of N fertilizers can result in the production of toxic metabolites
(nitrosamines, etc) that can have not only depressing effects on most soil micro-
organisms but also cause teratogenic, carcinogenic and allergic effects in higher
organisms (plants, animals and humans) through the food chain. In general the
dynamics of microbial population during the growing season for cacao seedlings on the
field, apart from effects of fertilizer types and rates can be attributed to the effects of
climatic factors and the excretory function of the root and moisture regimes during the
growing season (Bolton et al. 1992). The fertilizer treatments enhanced height of cacao
seedlings than control in both locations at 3, 4 and 5 months after transplanting (Table
6). Cacao plants were taller at Ibadan than Owena at 5 MAT. The differences in height
between the both locations were attributed to the differences in the nutrient- releasing
pattern and growing environmental conditions between the two locations. The
significant increases in the growth parameters of cacao seedlings at 3, 4 and 5 MAT in
Ibadan and Owena relative to the control under the fertilizer treatments can be
attributed to the nutrient contents of organic fertilizers applied which enhanced cacao
seedlings growth. This finding that the manure (GD, OMF and OF) improved the growth
of cacao is consistent with earlier findings of Adeniyan and Ojeniyi (2005) and Moyin-
Jesu (2007) who reported that organic manures supported the growth and development
of maize and coffee. Poor growth of cacao seedlings as a result of low nutrient status of
soil was generally observed in the unfertilized treatment plots in the two locations in
2012 indicating that the soils of both locations were low in fertility and not supportive
of good cacao growth. The higher plant height observed in Ibadan 2012 experiment at
3, 4 and 5 MAT, where the highest rate of NPK was applied (600 kg/ha) could be as a
result of the rapid release of nutrients following mineralization and the consequent
absorption for cacao growth. The number of branches recorded at control was lower
but not significantly different from plots where the fertilizer types were applied in
Ibadan and Owena at 3 MAT (Table 8). The similar performance of cacao seedlings with
regards to number of branches under control plots (Ibadan and Owena 3 MAT)
compared to organic fertilizer treatments might be due to the high initial nutrient
contents of the soils.
The values of growth parameters (plant height, number of leaves and stem diameter) at
15, 16 and 17 MAT due to manures of both animal and plant origins were higher
compared to that of inorganic origin (NPK 15: 15: 15 fertilizer, tables 9, 10 and 11) .
This might be due to presence of other vital nutrient elements like Ca, Mg, OC and other
micro-nutrients that are required for cacao seedlings growth which are absent in the
NPK 15: 15:15 fertilizer.(Ipinmoroti et al. 2002) The relatively taller plant under 600
kg/ha OF in Ibadan at 15, 16 and 17 MAT could have stemmed from the nutrient
contents of the organic fertilizers (essential soil nutrients) which though released
slowly, last longer in the soil for optimum crop performance (Lombin 1981). Titiloye et
al. (1985) have reported a survey of 45 waste materials which were found to be rich in
the nutrient elements (N, P, K, Ca, Mg, Zn, Cu, Fe and Mn contents). The farm wastes
therefore represent a potential source of nutrients that could be harnessed to boost
crop growth and productivity (Solomon and Ogeh 1995).
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Table 1: Physical and chemical characteristics of the soils before planting in

Ibadan and Owena (2012 experiments)
Soil Properties Ibadan 2012 Owena 2012
Experiment Experiment
Sand (%) 75.1 74.1
Silt ( Silt (%) 17.4 16.3
Clay (%) 7.5 9.6
Textural class Sandy loam Sandy loam
pH (water) 4.56 5.70
Organic carbon 0.43 1.51
(g/kg)
Organ Organic 1.26 1.61
matter (%)
Total I Nitrogen 1.26 0.14
(%)
Available P 8.56 13.52
(cmol/kg)
K+ (cmol/kg) 3.26 0.24
Ca Ca
++ ++ 6.00 2.60
(cmol/kg)
Mg++ (cmol/kg) 3.00 1.00
Na+ (cmol/kg) 2.52 0.15
Al+++ 1.21 1.86
H+(cmol/kg) 6.89 8.12
ECEC 19.88 13.97
Table 2: Chemical composition of the organic materials used

Properties Goat dung (GD) Organo-mineral Organic
fertilizer (OMF) fertilizer (OF)
pH (water) 8.17 7.00 7.30
Organic carbon (g/kg) 2.86 0.63 3.64
Organic matter (%) 4.80 1.09 6.27
Total nitrogen (%) 1.26 0.06 2.16
Available P (cmol/kg) 113.24 138.06 7.08
K (cmol/kg)
+ 0.41 0.19 5.56
Mg++ (cmol/kg) 1.20 1.00 6.00
Ca++ (c Ca++ cmol/kg) 2.60 2.00 13.10
Na+ (cmol/kg) 0.38 0.18 2.30
C:N 1:4 1:1 1:6
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Table 3: Effects of organic and inorganic fertilizer types and rates on albuscular
mycchorizal spore count (Ibadan and Owena 2012 experiments)
Treatments Ibadan 2012 Owena 2012
GD 600 28.00de 35.00bc
GD 400 47.33ab 54. 00a
GD 200 42.00abc 35.00ab
Control 26.87de 33.65bcd
OMF 600 37.66bcd 47.33a
OMF 400 20.00e 31.33bcd
OMF 200 41.00abcd 36.33ab
OF 600 28.33cde 44.66ab
OF 400 52.00a 51.33a
OF 200 38.33bcd 36.33ab
NPK 600 36.00bcd 34.00bc
NPK 400 33.66cd 33.00bcd

NPK 200 30.33cde 32.66bcd
Treatment means within each column followed by the same letters are not significantly
different from each other using Tukey’s HSD at 5% level
Table 4: Total population of nematodes as affected by fertilizer types and rates 15

MAT at Ibadan and Owena 2012 experiments
Fertilizers Rate (kg/ha) Ibadan Owena
Types 2012 2012
GD 600 63.99j 129.66b
400 252.31b 98.32f
200 244.32c 102.99d
Control 355.87a 195.43a
OMF 600 52.31k 81.32h
400 83.32h 117.65c
200 88.65g 55.65m
Control 365.67a 194.00a
OF 600 105.64f 89.32g
400 50.98l 101.98e
200 165.65d 72.31j
Control 357.12a 193.23a
NPK 600 14.00m 65.32k
400 138.64e 73.97i
200 80.31i 64.98i
Control 358.31a 195.32a
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different from each other using Tukey’s HSD at 5% level.
Table 5: Effects of fertilizer types and rates on colonies of bacteria, yeasts and
moulds in Ibadan and Owena (2012 experiments).
Ibadan 2012 Owena 2012
Bacteria Yeasts Moulds Bacteria Yeasts Moulds
(cfu/g) (cfu/g) (cfu/g) (cfu/g) (cfu/g) (cfu/g)
GD 600 183.33b 100.00bc 25.00c 38.33e 30.66c 10.66bc
GD 400 200.00b 75.00de 20.00cd 121.00b 60.00ab 10.00bc
GD 200 295.00a 120.00b 26.66c 71.00cd 65.00a 11.00bc

Control 71.43d 44.32g 15.43cd 31.54e 27.54cd 9.00c
OMF 600 196.67b 100.00bc 25.00c 60.00d 55.00ab 15.66bc

OMF 400 196.00b 203.33a 19.66cd 134.66b 55.66ab 9.00bc
OMF 200 100.00cd 95.00cd 22.00cd 100.00c 55.66.ab 20.66b
Control 68.76d 40.99g 16.98ed 30.76e 26.76cd 7.99c
OF 600 170.00b 190.00a 16.00ed 42.00e 56.66ab 8.66c
OF 400 120.00c 70.00e 20.00cd 92.00c 50.00b 50.00a
OF 200 176.67b 50.00ef 24.00c 64.66d 30.66c 9.33bc

Control 70.98d 42.23g 16.12ed 30.11e 25.32cd 7.89c
NPK 600 160.00b 54.67ef 37.33b 270.00a 25.00cd 10.66bc
NPK 400 176.67b 61.67ef 22.00cd 30.00e 20.00d 6.00c
NPK 200 180.00b 110.00cd 45.33a 35.33e 50.00b 16.66bc

CONTR 70.00d 43.33fg 15.00ed 30.00e 25.00cd 8.00c
Table 6: Effects of organic and inorganic fertilizer types and rates on plant height of cacao
seedlings in Ibadan and Owena (2012 Experiments)
Treatments Ibadan Owena
Months after transplanting (MAT) Months after transplanting (MAT)
Fertilizers Rates 3 4 5 3 4 5
(kg/ha)
Goat 600 40.20ab 44.80abc 5066abc 35.89ab 45.18abc 51.23ab
Dung .
400 42.83ab 49.50ab 53.33ab 36.22ab 53.97ab 63.09a
200 38.86ab 43.14bc 41.33bc 34.89ab 47.55abc 54.24ab
Control 34.89b 37.32c 38.12c 23.99b 34.67c 32.41c
Organo- 600 43.70ab 49.27ab 52.33ab 30.33ab 42.09abc 46.95abc
Mineral 400 46.40ab 51.67ab 55.00a 25.67b 42.32abc 46.88abc
Fertilizer 200 43.70ab 50.17ab 54.33ab 33.00ab 53.11ab 58.58ab
Control 35.43b 37.87c 37.98c 24.99b 34.56c 32.43c
Organic 600 43.33ab 49.41ab 54.33ab 31.28ab 43.56abc 46.04abc
manure 400 40.97ab 45.50abc 48.67abc 32.22ab 48.01abc 53.61ab
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200 42.53ab 47.35abc 52.33ab 39.53a 57.63a 62.52a

Control 36.00b 36.99c 39.00c 25.00b 35.00c 31.98c
NPK 600 48.07a 55.21a 56.00a 33.22ab 51.08ab 51.76ab
15:15:15 400 47.23ab 50.82ab 53.00ab 32.61ab 44.39abc 45.88abc
200 44.47ab 51.62ab 52.66ab 28.55ab 40.72bc 42.69bc
Control 35.84b 37.47c 38.66c 24.89b 34.75c 32.41c
different from each other using Turkey’s HSD at 5% level.
Table 7: Effects of organic and inorganic fertilizer types and rates on number of leaves of
cacao seedlings in Ibadan and Owena (2012 experiments)
Months After Transplanting Months After Transplanting (MAT)
(kg/ha)
Goat 600 14.50a 17.17abc 20.00ab 17.00ab 20.17ab 26.35a
Dung 400 12.60ab 14.47bcde 17.00bcd 17.79ab 20.47a 25.67a
200 18.83a 22.88a 25.67a 18.00a 19.13ab 21.77abc
Control 7.45bc 8.98c 9.65e 11.11ab 12.43b 13.00d
Organo- 600 15.60a 19.64ab 21.67ab 14.22ab 17.06ab 22.73abc
Mineral 400 15.30a 19.11ab 21.67ab 9.33b 13.12ab 15.70cd
Fertilizer 200 15.60a 19.11ab 21.67ab 14.89ab 19.55ab 26.40a
Control 7.89c 7.99e 10.00e 10.99ab 12.98b 13.11d
Organic 600 12.60ab 16.50abcd 19.00abc 14.33ab 16.81ab 23.76ab
manure 400 13.93a 17.73ab 21.33ab 15.66ab 20.29a 25.80a
200 15.70a 19.23ab 22.33ab 14.54ab 18.17ab 23.68ab
Control 6.99bc 7.98e 9.95e 11.00ab 12.55b 12.56d
NPK 600 6.43c 10.33cde 11.67cde 14.33ab 17.42ab 19.61abcd
15:15:15 400 6.53c 9.87de 11.33de 14.05ab 13.75ab 16.54bcd
200 13.03ab 16.33abcd 17.33bcd 10.89ab 16.65ab 15.27cd
Control 7.50bc 8.33e 9.67e 11.03ab 12.43b 12.89d
Table 8: Effects of organic and inorganic fertilizer types and rates on number of
branches of cacao seedlings in Ibadan and Owena (2012 experiments)
Months after transplanting Months after transplanting (MAT)
(kg/ha)
Goat 600 1.50a 2.06ab 2.38ab 0.00a 1.22ab 2.24ab
Dung 400 1.00a 1.85ab 2.53ab 0.00a 0.53abc 1.24ab
200 1.00a 1.84ab 2.15ab 0.22a 0.42abc 0.63ab
Control 0.23a 0.34b 0.56b 0.00a 0.00c 0.17b
Organo- 600 1.00a 2.15ab 2.48ab 0.00a 0.12c 1.18ab
Mineral 400 1.67a 2.59ab 3.60ab 0.00a 0.17c 0.63ab
Fertilizer 200 1.00a 1.66ab 2.07ab 0.53a 0.78abc 1.52ab
Control 0.24a 0.40b 0.60b 0.00a 0.00c 0.19b
Organic 600 1.33a 2.36ab 2.97ab 0.00a 1.31a 2.27a
fertilizer 400 2.33a 3.68a 4.10a 0.00a 0.83c 1.72ab
200 2.33a 3.75a 4.17a 0.44a 0.86abc 1.43ab
Control 0.27a 0.43b 0.76b 0.00a 0.01c 0.19b
NPK 600 1.00a 2.02ab 2.15ab 0.22a 0.61abc 1.05ab
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15:15:15 400 0.90a 1.80ab 2.03ab 0.00a 0.00c 0.59ab

200 2.17a 3.47a 3.57ab 0.00a 0.22bc 0.44b
Control 0.25a 0.34b 0.52b 0.00a 0.00c 0.18b
Table 9: Effects of organic and inorganic fertilizer types and rates on plant height of cacao
seedlings in Ibadan and Owena (2012 experiments)
Rates Months After Transplanting (MAT) Months After Transplanting (MAT)
(kg/ha) 15 16 17 15 16 17
Goat 600 71.67abc 73.00abcd 80.33abc 157.00ab 196.00ab 205.67a
Dung 400 67.67abc 69.00abcd 72.33bcd 159.67ab 192.33abc 196.33ab
200 77.67abc 81.00abcd 86.67abc 186.67a 191.33abc 192.67ab
Control 45.43c 47.43d 50.87d 108.32b 87.54f 90.65e
Organo- 600 70.33abc 77.67abcd 85.67abc 160.67ab 205.67a 207.33a
Mineral 400 86.33ab 91.33ab 95.33ab 142.00ab 137.33bcd 140.67bcde
Fertilizer 200 78.33abc 83.33abc 91.67abc 129.67ab 171.67abcd 175.00abc
Control 44.76c 47.76d 51,98d 104.98b 86.21f 88.00e
Organic 600 93.33a 99.67a 105.67a 152.67ab 179.00abc 184.33ab
Manure 400 84.67ab 90.33abc 96.67ab 149.00ab 161.00abcd 165.00abcd
200 87.67ab 90.33abc 96.67ab 146.67ab 160.33abcd 163.67abcd
Control 47.98c 48.56d 50.78d 105.98b 84.98f 85.99e
NPK 600 67.00abc 70.00abcd 80.00abc 113.50b 130.00cdef 136.00bcde
15:15:15 400 54.73bc 56.33cd 63.33cd 111.00b 137.33bcde 144.00abcd
200 69.33abc 73.67abcd 78.67abc 139.67ab 111.00def 115.33cde
Control 46.00c 48.00d 50.00d 106.00b 85.33f 87.00e
Table 10: Effects of organic and inorganic fertilizer types and rates on number of leaves
of cacao seedlings in Ibadan and Owena (2012 experiments)
Months After Transplanting (MAT) Months After Transplanting (MAT)
Rates
Fertilizers (kg/ha) 15 16 17 15 16 17
Goat 600 93.33a 93.33a 101.00a 77.00ab 101.33b 106.00b
Dung 400 42.67bc 43.67bcd 47.67bc 57.67abc 135.00ab 137.67ab
200 44.00bc 48.33bcd 53.33bc 83.00a 156.67ab 161.00ab
Control 21.33c 23.21d 24.54c 24.87bc 76.65b 82.00b
Organo- 600 60.00b 61.00abc 69.33ab 67.33abc 171.67ab 176.00ab
Mineral 400 45.67bc 47.33bcd 51.67bc 40.67abc 70.00b 72.33b
Fertilizer 200 32.67bc 35.67bcd 39.67bc 93.00a 306.00a 310.00a
Control 19.99c 22.98d 23.67c 24.98bc 79.09b 87.09b
Organic 600 51.33bc 58.00bcd 65.67b 56.67abc 114.33ab 121.00ab
Manure 400 33.67bc 35.33bcd 44.67bc 47.67abc 110.33ab 111.67ab
200 34.67bc 38.33bcd 43.67bc 38.33abc 114.33ab 118.00ab
Control 20.87c 24.98d 23.78c 25.00bc 79.78b 80.98b
NPK 600 60.00b 65.00ab 70.00ab 35.50abc 120.00ab 125.00ab
15:15:15 400 26.00c 29.67cd 37.33bc 50.67abc 48.00b 52.00b
200 37.00bc 40.00bcd 48.67bc 22.00bc 80.67b 83.00b
Control 20.33c 23.33d 24.67c 24.33bc 78.67b 81.00b
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Table 11: Effects of organic and inorganic fertilizer types and rates on number of
branches of cacao seedlings at 15, 16 and 17 MAT in Ibadan and Owena (2012
experiments)
Months After Transplanting (MAT) Months After Transplanting (MAT)
Rates
Fertilizers (kg/ha) 15 16 17 15 16 17
Goat Dung 600 12.86a 13.33a 15.33a 11.16ab 17.33abc 17.33abcd
400 9.33abc 10.00ab 12.66ab 8.00abc 24.00abc 26.33abcd
200 5.33abc 7.00ab 7.83ab 7.66abc 30.33ab 34.33a
Control 2.43c 3.55b 3.56b 3.51c 5.11c 6.56d
Organo- 600 7.00abc 8.00ab 7.00ab 12.06a 30.00ab 31.00abc
Mineral 400 6.33abc 8.00ab 8.00ab 6.00abc 13.66abc 16.66abcd
Fertilizer 200 5.00abc 5.66ab 7.00ab 9.50abc 25.66abc 27.66abcd
Control 2.33c 3.56b 3.67b 3.53c 4.99c 6.65d
Organic 600 9.66abc 11.66ab 12.66ab 9.16abc 24.66abc 33.00ab
Fertilizer 400 6.86abc 7.66ab 9.00ab 6.50abc 34.00a 38.00a
200 9.66abc 11.33ab 13.33ab 7.16abc 30.66ab 35.00a
Control 2.33c 3.76b 3.32b 3.56c 5.00c 6.67d
NPK 600 12.00ab 12.50ab 13.50ab 6.75abc 12.50abc 11.00bcd
15:15:15 400 3.53bc 3.90ab 3.90ab 6.00abc 9.00bc 10.66bcd
200 9.20abc 9.66ab 13.33ab 5.66abc 11.66abc 9.66cd
Control 2.33c 3.66b 3.66b 3.50c 5.00c 6.66d
CONCLUSION
This research work explored the effects of Goat Dung (GD), Sunshine Organic and
Organo-Mineral Fertilizers (OF and OMF) and NPK 15:15:15 on soil biological
properties and growth of cacao between 2012 and 2014 in two cacao growing ecologies
of south western, Nigeria. The results showed that organic manures applied increased
the abundance of Verscular Abuscular Mychorizal Spores (VAM) in the soil relative to
NPK and the control in Ibadan and Owena. Soil application of 400 kg/ha OF enhanced
AMF compared with other fertilizer types and the control in Ibadan. Application of 600
kg/ha NPK and 200 kg/ha OMF produced the lowest nematode count in Ibadan and
Owena. Similar trends were observed in Ibadan and Owena, where highest nematode
populations were recorded for the control plots.
The total count of soil bacteria was highest for 200 kg/ha GD and 600 kg/ha NPK at
Ibadan and Owena. Fungal populations at Ibadan and Owena were significantly
enhanced when 200 kg/ha NPK and 400 kg/ha OF were applied. Lower rates of
application of NPK fertilizer at 200 kg/ha led to increased fungal populations at both
Ibadan and Owena. In Ibadan (2012 experiment), application of NPK at 600 kg/ha gave
significant (P<0.05) plant height at both 3,4 and 5 MAT of cacao relative to the control
while 600 kg/ha OF gave significant (P<0.0.5) higher plant higher at both 15, 16 and 17
MAT relative to the control. Again, in Owena, 200 kg/ha OF enhanced plant height at 3, 4
and 5 MAT compared with the control. However, 200 kg GD enhanced higher plant
height at 15 MAT compared with the control and 600 kg/ha OMF significantly increased
height of cacao at 16 and 17 MAT respectively. Similar trends were obtained in Owena
ISSN 2320-1355
at application of 200 kg/ha OMF produced significantly higher number of leaves at 15,
16 and 17 MAT over the control. The effect of 200 kg/ha GD was significant on number
of leaves in Ibadan at 3, 4 and 5 MAT. Also, 600 kg/ha GD produced marked effect on
number of cacao leaves relative to the control at 15, 16 and 17 MAT.
Application of 400 and 200 kg/ha OF had profound effect on number of branches in
Ibadan at 3, 4 and 5 MAT compared with other treatments and the control. 600 kg/ha
GD had better influence on number of branches in Ibadan at 15, 16 and 17 MAT
compared with other treatments and the control.
In Owena at 4 and 5 MAT, the number of branches produced under 600 kg/ha OF was
higher compared with other treatments and the control. At 15 MAT, the number of
branches produced under 600 kg/ha OMF was greater than what obtained under other
treatments and the control followed by 600 kg/ha GD, while application of 400 kg/ha
OF produced higher number of branches at 16 and 17 MAT which was about 85 and 82
% increases over the control respectively. In order to suppress nematode population at
Ibadan, 600 kg/ha NPK is recommended. Adoption of application of 400 kg/ha OF and
200 kg/ha OMF is recommended for reducing soil nematode population at Owena.
Higher rate of GD (600 kg/ha) is recommended to promote bacterial population. 200
kg/ha OMF is recommended for increase in yeast population and 400 kg/ha OF for fungi
at Ibadan, while 400 kg/ha NPK is recommended for optimum fungi population at
Owena. In order to increase soil bacteria and yeast population at Ibadan and Owena to
attain rapid decomposition of organic matter, 200 kg/ha GD is recommended. The
fertilizer treatments enhanced the growth parameters, soil and leaf chemical
composition of cacao seedlings than the control in both locations, organic manures
performed better than the organo-minerals (OMF), while the organo-mineral fertilizers
were better than NPK.
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Journal of Global Biosciences

Soil and Pest Management,

MATERIALS AND METHODS

Soil samples collection and analysis

Exchangeable K, Ca, Mg and Na were determined by extraction with ammonium acetate

RESULTS AND DISCUSSION

organic fertilizer application, either as sole or in combination with inorganic fertilizers

Table 1: Physical and chemical characteristics of the soils before planting in

Table 2: Chemical composition of the organic materials used

NPK 400 33.66cd 33.00bcd

Control 27.33de 33.66bcd

Table 4: Total population of nematodes as affected by fertilizer types and rates 15

GD 600 183.33b 100.00bc 25.00c 38.33e 30.66c 10.66bc

GD 400 200.00b 75.00de 20.00cd 121.00b 60.00ab 10.00bc

GD 200 295.00a 120.00b 26.66c 71.00cd 65.00a 11.00bc

OMF 600 196.67b 100.00bc 25.00c 60.00d 55.00ab 15.66bc

OF 200 176.67b 50.00ef 24.00c 64.66d 30.66c 9.33bc

NPK 400 176.67b 61.67ef 22.00cd 30.00e 20.00d 6.00c

NPK 200 180.00b 110.00cd 45.33a 35.33e 50.00b 16.66bc

200 42.53ab 47.35abc 52.33ab 39.53a 57.63a 62.52a

15:15:15 400 0.90a 1.80ab 2.03ab 0.00a 0.00c 0.59ab

Whitehead AG, Hemming JR. 1965. A comparison of some quantitative methods of

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